Barbary lion

Characteristics

Barbary lion zoological specimens range in colour from light to dark tawny. Male lion skins had short manes, light manes, dark manes or long manes. Head-to-tail length of stuffed males in zoological collections varies from 2.35 to 2.8 m (7 ft 8 ¹⁄₂ in to 9 ft 2 in), and of females around 2.5 m (8 ft 2 in). Skull size varied from 30.85 to 37.23 cm (12 ⁵⁄₃₂ to 14 ²¹⁄₃₂ in). Some manes extended over the shoulder and under the belly to the elbows. The mane hair was 8 to 22 cm (3 to 8 ¹⁄₂ in) long.

In 19th century hunter accounts, the Barbary lion was claimed to be the largest lion, with a weight of wild males ranging from 270 to 300 kg (600 to 660 lb). Yet, the accuracy of such data measured in the field is questionable. Captive Barbary lions were much smaller but kept under so poor conditions that they might not have attained their full potential size and weight.

The colour and size of lions' manes was long thought to be a sufficiently distinct morphological characteristic to accord a subspecific status to lion populations. Mane development varies with age and between individuals from different regions, and is therefore not a sufficient characteristic for subspecific identification. The size of manes is not regarded as evidence for Barbary lions' ancestry. Instead, results of mitochondrial DNA research support the genetic distinctness of Barbary lions in a unique haplotype found in museum specimens that is thought to be of Barbary lion descent. The presence of this haplotype is considered a reliable molecular marker to identify captive Barbary lions. Barbary lions may have developed long-haired manes, because of lower temperatures in the Atlas Mountains than in other African regions, particularly in winter. Results of a long-term study on lions in Serengeti National Park indicate that ambient temperature, nutrition and the level of testosterone influence the colour and size of lion manes.

Taxonomic history

Felis leo was the scientific name proposed by Carl Linnaeus in 1758 for a lion type specimen from Constantine, Algeria. Following Linnaeus's description, several lion zoological specimens from North Africa were described and proposed as subspecies in the 19th century:

• Felis leo barbaricus described by the Austrian zoologist Johann Nepomuk Meyer in 1826 was a lion skin from the Barbary Coast.
• Felis leo nubicus described by Henri Marie Ducrotay de Blainville in 1843 was a male lion from Nubia that had been sent by Antoine Clot from Cairo to Paris and died in the Ménagerie du Jardin des Plantes in 1841.

In the 20th century, there has been much debate and controversy among zoologists on lion classification and validity of proposed subspecies:

• In 1939, Glover Morrill Allen considered F. l. barbaricus and nubicus synonymous with F. l. leo.
• Reginald Innes Pocock subordinated the lion to the genus Panthera, when he wrote about the Asiatic lion.
• In 1951, John Ellerman and Terence Morrison-Scott recognized only two lion subspecies in the Palearctic realm, namely the African lion Panthera leo leo and the Asiatic lion P. l. persica.
• Some authors considered P. l. nubicus a valid subspecies and synonymous with P. l. massaica.
• In 2005, P. l. barbarica, nubica and somaliensis were subsumed under P. l. leo.
• In 2016, IUCN Red List assessors used P. l. leo for all lion populations in Africa.

In 2017, the Cat Classification Task Force of the Cat Specialist Group subsumed the lion populations in North, West and Central Africa and Asia to P. l. leo.

Genetic research

Results of a phylogeographic analysis using samples from African and Asiatic lions was published in 2006. One of the African samples was a vertebra from the National Museum of Natural History (France) that originated in the Nubian part of Sudan. In terms of mitochondrial DNA, it grouped with lion skull samples from the Central African Republic, Ethiopia and the northern part of the Democratic Republic of the Congo.

While the historical Barbary lion was morphologically distinct, its genetic uniqueness remained questionable. In a comprehensive study about the evolution of lions in 2008, 357 samples of wild and captive lions from Africa and India were examined. Results showed that four captive lions from Morocco did not exhibit any unique genetic characteristic, but shared mitochondrial haplotypes with lion samples from West and Central Africa. They were all part of a major mtDNA grouping that also included Asiatic lion samples. Results provided evidence for the hypothesis that this group developed in East Africa, and about 118,000 years ago traveled north and west in the first wave of lion expansion. It broke up within Africa, and later in West Asia. Lions in Africa probably constitute a single population that interbred during several waves of migration since the Late Pleistocene. Genome-wide data of a wild-born historical lion specimen from Sudan clustered with P. l. leo in mtDNA-based phylogenies, but with a high affinity to P. l. melanochaita.

Former distribution and habitat

Historical accounts indicate that in Egypt lions occurred in the Sinai Peninsula, along the Nile, in the Eastern and Western Deserts, in the region of Wadi El Natrun and along the maritime coast of the Mediterranean. In the 14th century BC, Thutmose IV hunted lions in the hills near Memphis. The growth of civilizations along the Nile and in the Sinai Peninsula by the beginning of the second millennium BC and desertification contributed to isolating lion populations in North Africa.

Historical sighting and hunting records from the 19th and 20th centuries show that lions inhabited the range countries of the Atlas Mountains from Tunisia to Morocco.

In Libya, the Barbary lion persisted along the Mediterranean coast until the beginning of the 18th century, and was extirpated in Tunisia by 1890.

In Algeria, the Barbary lion occurred in the forested hills and mountains between the Pic de Taza in the east, Ouarsenis in the west and the Chelif River plains in the north. Lions also inhabited the forests and wooded hills of the Constantine Province and south into the Aurès Mountains. In the 1830s, lions may have already been eliminated along the coast and near human settlements. By the mid-19th century, the lion population had massively declined, since bounties were paid for shooting lions. The cedar forests of Chelia and neighbouring mountains harboured lions until about 1884. They disappeared in the Bône region by 1890, in the Khroumire and Souk Ahras regions by 1891, and in Batna Province by 1893. The last known sighting of a lion in Algeria occurred in 1956 in Beni Ourtilane District.

In Morocco, the last recorded shooting of a wild Barbary lion took place in 1942 near Tizi n'Tichka in the Atlas Mountains. A small remnant population may have survived in remote montane areas into the early 1960s.

Behaviour and ecology

In the early 20th century, when Barbary lions were not common anymore, they were sighted in pairs or in small family groups comprising a male and female lion with one or two cubs. Between 1839 and 1942, sightings of wild lions involved solitary animals, pairs and family units. Analysis of these sightings indicate that lions retained living in prides even when under increasing persecution, particularly in the eastern Maghreb. The size of prides was likely similar to prides living in sub-Saharan habitats, whereas the density of the Barbary lion population is considered to have been lower than in moister habitats.

When Barbary stags and gazelles became scarce in the Atlas Mountains, lions preyed on herds of livestock that were rather carefully tended. They also preyed on wild boar and red deer.

Sympatric predators in this area included the African leopard and Atlas bear.

In captivity

The lions kept in the menagerie at the Tower of London in the Middle Ages were Barbary lions, as shown by DNA testing on two well-preserved skulls excavated at the Tower between 1936 and 1937. The skulls were radiocarbon-dated to around 1280–1385 and 1420−1480. In the 19th century and the early 20th century, lions were often kept in hotels and circus menageries. In 1835, the lions in the Tower of London were transferred to improved enclosures at the London Zoo on the orders of the Duke of Wellington.

The lions in the Rabat Zoo exhibited characteristics thought typical for the Barbary lion. Nobles and Berber people presented lions as gifts to the royal family of Morocco. When the family was forced into exile in 1953, the lions in Rabat, numbering 21 altogether, were transferred to two zoos in the region. 3 of these were shifted to a zoo in Casablanca, with the rest being shifted to Meknès. The lions at Meknès were moved back to the palace in 1955, but those at Casablanca never came back. In the late 1960s, new lion enclosures were built in Temara near Rabat. Results of a mtDNA research revealed in 2006 that a lion kept in the German Zoo Neuwied originated from this collection and is very likely a descendant of a Barbary lion. Five lion samples from this collection were not Barbary lions maternally. Nonetheless, genes of the Barbary lion are likely to be present in common European zoo lions, since this was one of the most frequently introduced subspecies. Many lions in European and American zoos, which are managed without subspecies classification, are most likely descendants of Barbary lions. Several researchers and zoos supported the development of a studbook of lions directly descended from the King of Morocco's collection.

At the beginning of the 21st century, the Addis Ababa Zoo kept 16 adult lions. With their dark, brown manes extending through the front legs, they looked like Barbary or Cape lions. Their ancestors were caught in southwestern Ethiopia as part of a zoological collection for Emperor Haile Selassie of Ethiopia.

In March 2010, two lion cubs were moved to the Texas Zoo in Victoria, Texas, where efforts were made to preserve Barbary lions under the WildLink International conservation programme. Whether the cubs are of Barbary lion descent was not determined. In 2011, the Port Lympne Animal Park in Kent received a Barbary lioness as a mate for the resident male. As of June 2016, Wisconsin Big Cat Rescue in Rock Springs, Wisconsin, has two female lions born in 2001 that have been shown by DNA testing to be Atlas lions. The Living Treasures Wild Animal Park in New Castle, Pennsylvania, claims to keep a pair of Barbary lions in the park's collection. The Zoo des Sables d'Olonne, Vendee, France, also claims to have a male and female Atlas lion.

Cultural significance

The lion appeared frequently in early Egyptian art and literature. Statues and statuettes of lions found at Hierakonpolis and Koptos in Upper Egypt date to the Early Dynastic Period. The early Egyptian deity Mehit was depicted with a lion head. In Ancient Egypt, the lion-headed deity Sekhmet was venerated as protector of the country. She represented destructive power, but was also regarded as protector against famine and disease. Lion-headed figures and amulets were excavated in tombs in the Aegean islands of Crete, Euboea, Rhodes, Paros and Chios. They are associated with Sekhmet and date to the early Iron Age between the 9th and 6th centuries BC. The remains of seven mostly subadult lions were excavated at the necropolis Umm El Qa'ab in a tomb of Hor-Aha, dated to the 31st century BC. In 2001, the skeleton of a mummified lion was found in the tomb of Maïa in a necropolis dedicated to Tutankhamun at Saqqara. It had probably lived and died in the Ptolemaic period, showed signs of malnutrition and had probably lived in captivity for many years.

In Roman North Africa, lions were regularly captured by experienced hunters for venatio spectacles in amphitheatres.